A related hypothesis (Hansen et al.,
2003) suggests that orientation anisotropies in real-world imagery (Hansen & Essock,
2004; Kiel & Cristobal,
2000) are normalized (Schwartz & Simoncelli,
2001; Wainwright,
1999) by anisotropically weighted contrast gain control mechanisms similar to those described empirically by (Foley,
1994; Meese & Holmes,
2007; Petrov, Carandini, & McKee,
2005; Ross & Speed,
1991). The anisotropy of content in typical natural scenes consists of a relatively constant horizontal contrast peak across spatial frequency, presumably largely from foreshortening and horizon lines, and a vertical peak which is more variable with spatial frequency and more dependent on coincidental image content (cf. Hansen & Essock,
2004; Kiel & Cristobal,
2000; Switkes, Mayer, & Sloan,
1978). A different pattern is seen in cortical neurophysiology. Cortical neurons that are tuned to horizontal or vertical orientations are more prevalent than those preferring oblique orientations, in several species including primates (Coppola, White, Fitzpatrick, & Purves,
1998; Kennedy, Martin, Orban, & Whitteridge,
1985; Mansfield,
1974; Mansfield & Ronner,
1978; Yu & Shou,
2000), and human evoked potentials and fMRI responses show a stronger response to horizontal and vertical gratings (Furmanski & Engel,
2000; Maffei & Campbell,
1970; Zemon, Gutowski, & Horton,
1983). Across spatial and temporal parameters, this bias is seen to be greatest in cells preferring higher spatial frequencies (and lower temporal rates) in the central visual field (Leventhal & Hirsch,
1977; Mansfield,
1974), similar to the human behavioral effect (Berkeley, Kitterle, & Watkins,
1975; Camisa, Blake, & Lema,
1977; Campbell, Kulikowski, & Levinson,
1967; Essock & Lehmkuhle,
1982). More recent studies indicate that this numerical bias is greater for horizontal than for vertical (Coppola & White,
2004; Li, Peterson, & Freeman,
2003). Thus, if one assumes that neurons tuned to similar spatial and temporal frequencies contribute most to a neuron's gain-control pool (Heeger,
1992), such a numeric anisotropy would predict a larger horizontal effect at higher spatial frequencies. One purpose of the present study was to assess the horizontal effect across spatial frequency in order to compare the findings to these two predictions: whether the anisotropy is found at all spatial frequencies (like the bias in scene content), mainly at high spatial frequencies (as with the physiological bias), or whether something altogether different, and unpredicted, occurs.